3. Marco teórico
3.3. Metodologías activas en Educación Infantil
A node-by-node description of all groups recovered in the combined analysis with CO coding for polymorphisms (fig. 37) is presented below. Included are the definition of the node, the nodal support as represented by jackknife (JK) resampling support and decay index (DI), and the unambiguous morphological and mo- lecular synapomorphies (appendices 4 and 5 list all morphological synapomorphies for the morphology-only and combined analyses, re- spectively). The effects of the TS coding for polymorphic characters and of exclusion of taxa without IRBP sequence are presented as changes in topology or in support in the morphology-only or combined analyses. Figure 40 summarize the pattern of recovery of the clades and their jackknife support in the different analyses.
Node 1
Composition: Sigmodontinae (sensu Reig, 1984).
Partitioned evidence: clade recovered in both morphology-only and IRBP-only analyses.
Morphological synapomorphies: furred tail (12), zygomatic plate anterior to M1 alveolus (29), derived hyoid pattern (48), first rib articulates with CE7 and TL1 (79), humerus without entepicondylar foramen (82), lateral bacular mounds present (85), and dorsal prostates present (95).
Molecular synapomorphies: 34, including 16 unique and unreversed.
TS coding: no change. Reduced analysis: no change.
Remarks: Monophyly of the subfamily Sig- modontinae sensu Reig (1984), i.e., excluding neotomines, peromyscines, and tylomyines, has been amply demonstrated in recent phylogenet- ic analyses using IRBP dataset (D’Elı´a, 2003; Weksler, 2003; Jansa and Weksler, 2004). Most of the morphological synapomorphies listed above for the subfamily were recognized in previous studies (Carleton, 1980; Voss, 1993; Steppan, 1995); the characters related to the tail and to the zygomatic plate, however, need to be reassessed in analyses with denser sampling. Node 2
Composition: Sigmodontinae minus Thomas- omys.
Nodal support: JK 5 78%, DI 5 4.
Partitioned evidence: clade recovered in both morphology-only and IRBP-only analyses.
Morphological synapomorphies: eight mam- mae (1), pes with squamate plantar surface (6), broad zygomatic plate (28), alisphenoid strut absent (38), capsular process of lower incisor alveolus absent (45), humerus with supratro- chlear foramen (83), and subequal proximal edge of trochlear process and posterior articular facet of calcaneum (84).
Molecular synapomorphies: 2, including 1 unique and unreversed.
TS coding: reduction of nodal support. Reduced analysis: slight reduction of nodal support.
Remarks: Monophyly of the thomasomyine group sensu Hershkovitz (1962; 1966a) (i.e., including Delomys and Thomasomys among others) was first contested by Voss (1993) and subsequently refuted by recent phyloge- netic analyses with dense taxon sampling (Smith and Patton, 1999; D’Elı´a, 2003; Weksler, 2003). Node 3
Composition: Wiedomys+ Oryzomyini. Nodal support: JK 5 77%, DI 5 4.
Partitioned evidence: clade recovered in the morphology-only analysis but not in the IRBP- only analysis.
Morphological synapomorphies: weakly cu- neate interorbit with small crests (22), medium palate (32), M2 without protoflexus (64), and 12 ribs (78).
Molecular synapomorphies: none. TS coding: reduction of nodal support. Reduced analysis: no change.
Remarks: The sister group relationship be- tween Wiedomys and oryzomyines was also recovered in the morphological analysis of Steppan (1995; see fig. 5B), which included a larger sample of other sigmodontine tribes. Molecular studies, however, strongly challenge these results (Smith and Patton, 1999; D’Elı´a, 2003; Weksler, 2003).
Node 4
Composition: Oryzomyini.
Nodal support: JK 5 98%, DI 5 5.
Partitioned evidence: clade recovered in both morphology-only and IRBP-only analyses.
Morphological synapomorphies: ungual tufts absent on D1, developed in remaining hindfoot digits (7), incisive foramina do not reach M1 (31), tegmen tympani absent (42), M1 with undivided anterocone (58), enamel connection between paracone and protocone at middle of protocone on M1 (61).
Molecular synapomorphies: no change can be unambiguously assigned to node 4 because Oryzomys hammondi, the first taxon to branch off within the Oryzomyini, lacks IRBP data. Eight molecular synapomorphies are found when ACCTRAN optimization is employed; the same synapomorphies are recovered in the reduced analysis.
TS coding: no change.
Reduced analysis: Decay index increases to 9. Remarks: Corroboration for oryzomyine monophyly sensu Voss and Carleton (1993) comes from analyses using morphological data (Steppan, 1995) and nuclear genes (Weksler, 2003). Studies employing cytochrome b, how- ever, fail to recover such group (Smith and Patton, 1999; Bonvicino and Moreira, 2001; Bonvicino et al., 2003; D’Elı´a, 2003). As discussed by Weksler (2003), these results are probably due to the saturation of the phyloge- netic signal in this rapid-evolving mitochondrial gene.
Node 5
Composition: Oryzomyini minus Oryzomys hammondi.
Nodal support: JK 5 28%, DI 5 1.
Partitioned evidence: none; O. hammondi is recovered as the sister group of Oecomys, well nested within oryzomyines, in the morphology- only analysis. O. hammondi does not have IRBP
data and consequently was not included in the IRBP-only and reduced analyses.
TS coding: O. hammondi is recovered within clade B* in both combined and morphology- only analyses.
Morphological synapomorphies: long palate (32), simple posterolateral palatal pits (34), reduced capsular process of lower incisor alveolus (45), and M3 without posteroloph (68). Remarks: The position of O. hammondi among oryzomyines is unarguably the least secure in the present analysis. The hypothesis of O. hammondi as the most basal oryzomyine should be viewed with caution because of the lack of strong support for the basal relation- ships of oryzomyines, and because of the contradictory results of the different analyses. Nevertheless, the basal position could explain why O. hammondi has always been regarded as an oryzomyine with obscure relationships (e.g., Hershkovitz, 1948; Hershkovitz, 1970; Musser and Carleton, 1993).
Node 6
Composition: Zygodontomys and Scolomys (clade A).
Nodal support: JK 5 53%, DI 5 1.
Partitioned evidence: clade recovered in the morphology-only analysis but not in the IRBP- only analysis.
Morphological synapomorphies: narrow in- terparietal (26) and absence of mesoloph on M3 (67).
Molecular synapomorphies: none.
TS coding: increased nodal support in the combined analysis; clade not recovered in the morphology-only analysis.
Reduced analysis: increased nodal support. Remarks: Scolomys has most of the morpho- logical synapomorphies that characterize the oryzomyines (Voss and Carleton, 1993; Go´ mez- Laverde et al., 2004), but morphological char- acters do not present evidence about its re- lationship within the tribe. Analyses of cyto- chrome b data recover Scolomys outside oryzomyines (Smith and Patton, 1999; D’Elı´a, 2003; but see Garcia, 1999), but the position of Scolomys as a basal oryzomyine is secured in both combined and IRBP-only analyses (see also Weksler, 2003). On the other hand, evidence for the clustering of Scolomys and Zygodontomys in a monophyletic group comes mostly from morphological data.
Node 7
Composition: Zygodontomys.
Nodal support: JK 5 100%, DI 5 34. Partitioned evidence: clade recovered in both morphology-only and IRBP-only analyses.
Morphological synapomorphies: tail slightly bicolored (11), strongly cuneate interorbit with overhanging crests (22), incisive foramina pass M1 (31), mental foramen located at diastema (44), masseteric crests reach anterior to m1 procingulum (47), m2 with 3 roots (51), proto- cone and paracone forming single dentine basin without enamel connection on M1 (61), M1 and M2 without mesolophs (62), m1 and m2 without mesolophids (73), and m3 without posteroflexid (77).
Molecular synapomorphies: 25, including 15 unique and unreversed.
TS coding: no change. Reduced analysis: no change.
Remarks: The placement of Zygodontomys as a basal oryzomyine is well secured by combined and IRBP-only results (see also Weksler, 2003). Corroboration of Zygodontomys monophyly is an expected result (cf. Voss, 1991; Bonvicino et al., 2003). Previously regarded as a subspecies of Z. brevicauda (see Voss, 1991), Z. cherriei displays an impressive number of morphologi- cal (7) and molecular (8) differences to Z. brevicauda and is considered here as distinct species.
Node 8
Composition: Oryzomyini minus O. ham- mondi, Zygodontomys, and Scolomys.
Nodal support: JK 5 49%, DI 5 1.
Partitioned evidence: clade recovered in the IRBP-only analysis but not in the morphology- only analysis.
Morphological synapomorphies: M2 with protoflexus (64).
Molecular synapomorphies: 5, including 2 unique and unreversed.
TS coding: O. hammondi is recovered within clade B* in both combined and morphology- only analyses.
Reduced analysis: increased nodal support. Remarks: Although morphological evidence for this ‘‘core oryzomyine’’ clade is meager, it receives high nodal support from the molecular data (see also Weksler, 2003).
Node 9
Composition: Handleyomys, Oecomys, and 11 species of Oryzomys belonging to 8 groups: alfaroi, melanotis (O. rostratus), chapmani, albigularis (O. albigularis and O. levipes), megacephalus, yunganus, talamancae, and niti- dus (O. lamia, O. macconnelli, and O. russatus). (clade B).
Nodal support: JK 5 58%, DI 5 1.
Partitioned evidence: clade recovered in the IRBP-only analysis but not in the morphology- only analysis. A similar clade is recovered in the
latter analysis, in which O. alfaroi, O. rostratus, and O. chapmani are excluded and O. hammondi is included.
Morphological synapomorphies: pes with smooth plantar surface (6), and flexi meet at midline on M1 (57).
Molecular synapomorphies: 4, including 2 unique.
TS coding: O. hammondi is also included in clade B* in the combined and morphology-only analyses. Amphinectomys is also included in clade B* in the morphology-only analysis.
Reduced analysis: increased nodal support. Remarks: Previous morphological, molecu- lar, and allozymic studies recovered results partially consistent with clade B. Phenetic morphological analyses cluster Oecomys, the nitidus and megacephalus groups, but not the albigularis group (Patton and Hafner, 1983). Patton and da Silva (1995) found a clade including Oecomys, Oryzomys yunganus, O. megacephalus, O. nitidus, and O. macconnelli using both weighted parsimony and distance analyses of cytochrome b. Finally, analysis of allozymic data (Dickerman and Yates, 1995) recovered a clade containing O. albigularis, O. nitidus, and O. megacephalus.
Node 10
Composition: Oryzomys nitidus species group (O. lamia, O. macconnelli, and O. russatus).
Nodal support: JK 5 94%, DI 5 3.
Partitioned evidence: clade recovered in both morphology-only and IRBP-only analyses.
Morphological synapomorphies: tail strongly bicolored (11), m1 with 3 roots (50), and anapophyses present on TL17 (80).
Molecular synapomorphies: 4, including 1 unique.
TS coding: clade recovered in the combined analysis and in some fundamental cladograms of the morphology-only analysis.
Reduced analysis: no change.
Remarks: The nitidus group of species has been recognized early in Oryzomys taxonomic history (Thomas, 1901), but only recently it has been shown as a monophyletic group (Weksler, 1996; Percequillo, 1998; Musser et al., 1998). Phylogenetic analyses based on cytochrome b data have failed to recover the group (Bonvicino and Moreira, 2001), probably due to phylogenetic signal saturation. The six species recognized for the group (table 2) display a distinctive set of characters relative to other Oryzomys species complexes, but only three unambiguous morphological synapomor- phies characterize the group, all of them homoplasious within the Oryzomyini. The
presence of three roots in the first lower molar is the least homoplasious among them, observed additionally in the Oryzomys albigularis group, Oryzomys angouya, Microryzomys, and Oe- comys trinitatis.
Node 11
Composition: Oryzomys lamia and O. russa- tus.
Nodal support: JK 5 100%, DI 5 9. Partitioned evidence: clade recovered in both morphology-only and IRBP-only analy- ses.
Morphological synapomorphies: developed capsular process of lower incisor alveolus (45), m1 without ectolophid (72).
Molecular synapomorphies: 9, including 5 unique and unreversed.
TS coding: no change. Reduced analysis: no change.
Remarks: O. lamia was considered a junior synonym of O. russatus by Musser et al. (1998), but the impressive number of morphological (6) and molecular (9) differences between the two species warrants the recognition of species status to each taxon, as proposed by Bonvicino et al. (1999).
Node 12
Composition: Handleyomys, Oecomys, and 8 species of Oryzomys belonging to 7 groups: alfaroi, melanotis (O. rostratus), chapmani, albigularis (O. albigularis and O. levipes), megacephalus, yunganus, and talamancae (clade B minus nitidus group).
Nodal support: JK 5 7%, DI 5 1. Partitioned evidence: none.
Morphological synapomorphies: none. Molecular synapomorphies: 2.
TS coding: clade recovered in some fundamental cladograms of combined analy- sis.
Reduced analysis: clade not recovered. Remarks: Neither morphology nor IRBP provides phylogenetic signal for a robust reso- lution of the basal structure of clade B. Nevertheless, the nitidus group is recovered as the basal group in clade B in most analyses. Additional characters are needed to formulate a solid phylogenetic hypothesis for members of this clade.
Node 13
Composition: Oecomys.
Nodal support: JK 5 95%, DI 5 4.
Partitioned evidence: clade recovered in both morphology-only and IRBP-only analyses.
Morphological synapomorphies: pes with extremely developed interdigital pads (5),
strongly cuneate interorbit with overhanging crests (22), parietal with deep lateral expansion (25), narrow zygomatic plate (28), M2 with accessory loph to paracone (65), M3 with posteroloph (68).
Molecular synapomorphies: 2. TS coding: no change. Reduced analysis: no change.
Remarks: Monophyly of Oecomys was sug- gested by chromosomal data (Gardner and Patton, 1976) and supported by analyses of mitochondrial genes (Patton and da Silva, 1995; Andrade and Bonvicino, 2003) and nuclear genes (Weksler, 2003). Oecomys is the only oryzomyine taxon with morphological speciali- zations for arboreal life, with several of them serving as synapomorphies for the genus. Node 14
Composition: Oecomys bicolor, Oe. trinitatis, Oe. mamorae, and Oe. concolor.
Nodal support: JK 5 67%, DI 5 1.
Partitioned evidence: clade recovered in the morphology-only analysis but not in the IRBP- only analysis.
Morphological synapomorphies: zygomatic plate at same level as M1 alveolus (29).
Molecular synapomorphies: none. TS coding: no change.
Reduced analysis: clade found in both combined and morphology-only analyses.
Remarks: The basal position of Oe. cathe- rinae within Oecomys needs to be substanti- ated by supplementary characters and addition- al analyses employing denser taxonomic sam- pling.
Node 15
Composition: Oecomys bicolor and Oe. trini- tatis.
Nodal support: JK 5 60%, DI 5 1.
Partitioned evidence: clade recovered in the morphology-only analysis but not in the IRBP- only analysis.
Morphological synapomorphies: lacrimal ar- ticulates mainly with maxillary (21) and de- veloped capsular process of lower incisor alveolus (45).
Molecular synapomorphies: none. TS coding: increased nodal support. Reduced analysis: no change. Node 16
Composition: Oecomys concolor and Oe. mamorae.
Nodal support: JK 5 98%, DI 5 5.
Partitioned evidence: clade recovered in both morphology-only and IRBP-only analy- ses.
Morphological synapomorphies: subtle coun- tershading (15) and derived carotid circulation pattern 2 (37).
Molecular synapomorphies: 2, including 1 unique and unreversed.
TS coding: slight decrease in nodal support. Reduced analysis: no change.
Remarks: The sister group relationship be- tween Oecomys concolor and Oe. mamorae is the only phylogenetic hypothesis within Oecomys strongly supported by both molecular and morphological data.
Node 17
Composition: Handleyomys and 8 species of Oryzomys belonging to 7 groups: alfaroi, melanotis (O. rostratus), chapmani, albigularis (O. albigularis and O. levipes), megacephalus, yunganus, and talamancae.
Nodal support: JK 5 12%, DI 5 1. Partitioned evidence: none.
Morphological synapomorphies: M3 with- out hypoflexus (or diminutive) (69) and sub- terminal flexure of vesicular gland irregularly lobed (97).
Molecular synapomorphies: none.
TS coding: clade recovered in the com- bined analysis and in some fundamental cladograms of the morphology-only analy- sis.
Reduced analysis: clade not recovered. Remarks: Neither morphology nor IRBP provides phylogenetic signal for a robust reso- lution of the internal structure of clade B. Additional characters are needed to formulate a solid phylogenetic hypothesis for members of this clade.
Node 18
Composition: Handleyomys, Oryzomys al- faroi, O. chapmani, and O. rostratus.
Nodal support: JK 5 78%, DI 5 4.
Partitioned evidence: clade recovered in the IRBP-only analysis but not in the morphology- only analysis.
Morphological synapomorphies: presence of labial accessory root on m1 (50), m2 with 3 roots (51), and M2 with two fossetti at mesoflexus position (66).
Molecular synapomorphies: 3.
TS coding: increased nodal support in combined analysis. In the morphology-only analysis, Handleyomys is found as the sister group of Amphinectomys; the two are then connected to the alfaroi-chapmani-rostratus clade.
Reduced analysis: no change.
Remarks: In the previous analysis of IRBP sequences, Handleyomys also appears as the
sister group to the alfaroi and melanotis species groups of Oryzomys in a highly supported clade (Weksler, 2003). Morphological data by itself does not corroborate the position of Handley- omys as the sister group of the alfaroi-melanotis- chapmani complex—in the CO morphology- only tree, Handleyomys appears as the sister group to the Oryzomys albigularis group. Nevertheless, the signal provided by IRBP data is not falsified by the morphological data; in fact, the clade including Handleyomys and the alfaroi-rostratus-chapmani groups has a series of compelling morphological synapomorphies in the combined tree; particularly, the number of roots of the second lower molars is unique among taxa within clade B.
Node 19
Composition: Oryzomys alfaroi, O. chapmani, and O. rostratus.
Nodal support: JK 5 90%, DI 5 4.
Partitioned evidence: clade recovered in the morphology-only analysis. O. chapmani lacks IRBP sequence data, but O. alfaroi and O. rostratus are recovered as sister groups in the IRBP-only analysis.
Morphological synapomorphies: large sphe- nopalatine vacuities (36), flexi not interpene- trating on M1 (57), and enamel bridge connec- tion between paracone and protocone at middle of protocone on M1 (61).
Molecular synapomorphies: 9, including 4 unique and unreversed.
TS coding: increased nodal support. Reduced analysis: no change.
Remarks: The three species groups, chap- mani, alfaroi, and melanotis, have long been considered to be associated (Goldman, 1918), but besides a previous IRBP analysis (Weksler, 2003), no phylogenetic evidence has been pre- viously presented for their close relationship. In the CO morphology-only tree, this clade is recovered within clade D*, but the TS analysis places it within clade B*.
Node 20
Composition: Oryzomys chapmani and O. rostratus.
Nodal support: JK 5 62%, DI 5 1.
Partitioned evidence: none; O. rostratus is recovered as the sister group of O. alfaroi in the morphology-only analysis. O. chapmani does not have IRBP data and consequently was not included in the IRBP-only and reduced analy- ses.
Morphological synapomorphies: m1 without ectolophid (72).
TS coding: increased nodal support in the combined analysis.
Remarks: Relationships within the alfaroi- melanotis-chapmani complex warrants further analyses with denser taxonomic sampling. The chapmani and alfaroi species groups have been considered to be close taxa since Merriam (1901) recognized the melanotis and chapmani species groups, with the latter including O. rhabdops (member of the alfaroi group), and Goldman (1918) recognized the melanotis and alfaroi groups, with the latter including O. chapmani and O. saturatior. Musser and Carle- ton (1993), however, asserted that the alfaroi group may be more closely related to the melanotis complex.
Node 21
Composition: Five species of Oryzomys belonging to 4 groups: albigularis (O. albigularis and O. levipes), megacephalus, yunganus, and talamancae.
Nodal support: JK 5 20%, DI 5 1. Partitioned evidence: none.
Morphological synapomorphies: tail slightly bicolored (11) and ampullary glands forming tufts of tubules (96)
Molecular synapomorphies: none.
TS coding: clade recovered in the combined analysis and in some fundamental cladograms of the morphology-only analysis.
Reduced analysis: clade not recovered. Node 22
Composition: Oryzomys megacephalus and O. yunganus.
Nodal support: JK 5 99%, DI 5 6.
Partitioned evidence: clade recovered in the IRBP-only analysis but not in the morphology- only analysis.
Morphological synapomorphies: derived ca- rotid circulation pattern 1 (37) and absence of capsular process of lower incisor alveolus (45).
Molecular synapomorphies: 5, including 2 unique and unreversed.
TS coding: clade recovered in the combined analysis and in some fundamental cladograms of the morphology-only analysis.
Reduced analysis: no change.
Remarks: The two morphological synapo- morphies observed for this clade are unique transformations among members of clade B. In particular, the carotid circulation pattern 1 displayed by species of the yunganus and megacephalus groups is observed only in a few other oryzomyines such as Oligoryzomys and in Neacomys musseri. The closeness of yunganus and megacephalus groups has been recognized before. For example, Musser et al. (1998: 323) stated that ‘‘[members] of the Oryzomys mega- cephalus and O. yunganus groups seem much
alike compared with the other [Oryzomys species groups]. Were it not for the distinction between the two provided by molar occlusal patterns, we would confidently include O. yunganus and O. tatei [5 yunganus group] along with O. megacephalus and O. laticeps [mega- cephalus group sensu Musser et al., 1998] in one group apart from the trans-Andean species, which seem to form a tight morphological cluster despite their striking external differences, and members of the O. nitidus group’’.
Node 23
Composition: Oryzomys talamancae, O. albi- gularis, and O. levipes.
Nodal support: JK 5 52%, DI 5 1. Partitioned evidence: none.
Morphological synapomorphies: reduced sphenopalatine vacuities (36), flexi deeply inter- penetrating on M1 (57), and anapophyses present on TL17 (80).
Molecular synapomorphies: 4, including 1 unique among oryzomyines.
TS coding: clade recovered in the combined analysis and in some fundamental cladograms of the morphology-only analysis.
Reduced analysis: clade not recovered. Remarks: O. talamancae is recovered as the sister group of the nitidus species group in the IRBP-only and reduced analyses, with moder- ate nodal support. Additional data are needed to resolve the conflict that results in a poorly supported hypothesis, although nodal support is higher in the TS combined analysis.
Node 24
Composition: Oryzomys albigularis species group (O. albigularis and O. levipes).
Nodal support: JK 5 96%, DI 5 6.
Partitioned evidence: clade recovered in the morphology-only analysis. O. levipes does not have IRBP data and consequently was not included in the IRBP-only and reduced analy- ses.
Morphological synapomorphies: amphoral interorbit with square edges (22), medium palate (32), m1 with 3 roots (50), M1 with deeply divided anterocone (58), and M3 with posteroloph (68).
TS coding: no change.
Remarks: The albigularis group has been recognized since the early classification of